Gravitropic reaction of primary seminal roots ofZea mays L. influenced by temperature and soil water potential

The growth of the primary seminal root of maize (Zea mays L.) is characterized by an initial negative gravitropic reaction and a later positive one that attains a plagiotropic liminal angle. The effects of temperature and water potential of the surrounding soil on these gravitropic reactions were studied. Temperatures of 32, 25, and 18C and soil water potentials of −5,−38, and −67 kPa were imposed and the direction of growth was measured for every 1 cm length of the root. The initial negative gravitropic reaction extended to a distance of about 10cm from the graln. Higher temperatures reduced the initial negative gravitropic reaction. Lower soil water potential induced a downward growth at root emergence. A mathematical model, in which it was assumed that the rate of the directional change of root growth was a sum of a time-dependent negative gravitropic reaction and an establishment of the liminal angle, adequately fitted the distance-angle relations. It was suggested that higher temperatures and/or a lower water potential accelerated the diminution of the intitial negative gravitropic reaction.     

Effect of soil water content on the gravitropic behavior of nodal roots in maize

The direction of root growth is an important factor that determines the spatial distribution of roots in the soil. The influence of soil water content on the direction of growth of maize nodal roots was studied both in the field and in the greenhouse. In the field experiment, the one plot was regularly irrigated (I-plot) and the other non-irrigated (N-plot). In the greenhouse experiment, three water treatments were conducted on plants grown in pots: continuously wet (CW), early drying (ED), and late drying (LD). The direction of root growth was quantified by the angle from the vertical, measured at 1 cm intervals for 10 cm from the first five internodes. Nodal roots grew more vertically in the N-plot and ED treatment than those in the I-plot and CW treatment. This was due to the decrease of the initial angle and/or the liminal angle. It is therefore thought that two events regulate the growth direction of nodal roots under dry soil conditions: gravitropic bending at root emergence from the stem and the later establishment of the angle of growth. Nodal roots appearing after rewatering in the ED treatment grew in a similar direction as those in the CW treatment. It follows from this that the water content of the surrounding soil has a direct effect on the direction of growth. Nodal roots that emerged in rapidly drying soil in the LD treatment ceased growing after showing negative gravitropism. The possible mechanisms determining the growth direction of nodal roots in drier soils are discussed.     

Plagiogravitropism of maize roots

The direction of root growth can be studied by analyzing the trajectories of roots growing in soil. Both the primary seminal root and nodal roots of maize attain a preferred, or liminal, angle of growth that deviates from the vertical. These roots are said to be plagiogravitropic. Experiments using plants grown in soil-filled boxes revealed that the primary seminal root is truly plagiogravitropic. It shows both positive and negative gravitropism in response to gravity stimuli and tends to maintain its direction even after growing around obstacles. These are experimental results suggesting that plagiogravitropic growth is controlled by internal factors. The orientation of the grain affects the establishment of the liminal angle of the primary seminal root, and both the position of their node of origin and the root diameter are closely related to the plagiogravitropic behaviour of nodal roots. Several external factors are also known to influence plagiogravitropism. Low soil water content causes a decrease in the angle of growth and soil mechanical resistance suppresses the gravitropic curvature. Plagiogravitropic behaviour of both seminal and nodal roots plays a significant role in shaping the root system.     

A no hydrotropic response root mutant that responds positively to gravitropism in Arabidopsis

For most plants survival depends upon the capacity of root tips to sense and move towards water and other nutrients in the soil. Because land plants cannot escape environmental stress they use developmental solutions to remodel themselves in order to better adapt to the new conditions. The primary site for perception of underground signals is the root cap (RC). Plant roots have positive hydrotropic response and modify their growth direction in search of water. Using a screening system with a water potential gradient, we isolated a no hydrotropic response (nhr) semi-dominant mutant of Arabidopsis that continued to grow downwardly into the medium with the lowest water potential contrary to the positive hydrotropic and negative gravitropic response seen in wild type-roots. The lack of hydrotropic response of nhr1 roots was confirmed in a system with a gradient in air moisture. The root gravitropic response of nhr1 seedlings was significantly faster in comparison with those of wild type. The frequency of the waving pattern in nhr1 roots was increased compared to those of wild type. nhr1 seedlings had abnormal root cap morphogenesis and reduced root growth sensitivity to abscisic acid (ABA) and the polar auxin transport inhibitor N-(1-naphtyl)phtalamic acid (NPA). These results showed that hydrotropism is amenable to genetic analysis and that an ABA signaling pathway participates in sensing water potential gradients through the root cap.     

Influence of temperature and water potential on root growth of white oak

Root growth of white oak ( Quercus alba L.) was observed under field conditions using a rhizotron. The effects of temperature, soil water potential, and leaf water potential were evaluated on three measures of root growth and development: root elongation rate, number of growing roots, and root growth intensity (sum of projected root area compared to the total root viewing area). Root elongation rate was linearly related to changes in soil temperature and soil water potential. At soil temperatures less than 17deg;C, temperature was the dominant factor affecting rate of growth, bat at temperatures greater than 17°C soil water potential became the important factor. Unlike root elongation rate, the number of growing roots and root growth intensity increased at cold soil temperatures (8°C) and at soil water potentials of-0.3 to -0.8 MPa. At high soil water potentials (-0.1 MPa) root elongation rate reached a maximum while the number of growing roots and root growth intensity were low. These differences showed that root growth and development were not exclusively affected by the soil environment. In addition, the relationship between root growth and predawn leaf water potential suggested that root growth was a contributing factor to the drought resistance of white oak.     

干旱区膜下滴灌条件下土壤深层水对棉花根系生长、分布及产量的影响

在土柱栽培条件下研究膜下滴灌土壤深层水对棉花根系生长的影响及与植株地上部生长的关系,设置土壤(60～120 cm)有深层水和无深层水2个处理,每处理设2个生育期间灌溉处理,分别为田间持水量70%和55%.结果表明：棉花总根质量密度、40～120 cm土层根长密度、根系活力等与地上部干质量间均具有显著的相关关系.生育期间耕层70%田间持水量条件下,土壤有深层水处理的总根质量密度与无深层水处理无明显差异,但40～120 cm土层的根长密度增加,根系活力增强,提高了土壤贮备水消耗量,增加了地上部干质量,最终获得较高的经济产量及水分利用效率.土壤有深层水条件下,生育期间耕层55%田间持水量处理的根冠比较大,40～120 cm土层根长密度和80～120 cm土层根系活力相对较高,土壤贮备水消耗量大幅提高,但仍无法弥补生育期间水分亏缺对根系及地上部生物量造成的负面影响,导致经济产量显著低于70%田间持水量处理.综上,充足的土壤深层水配合生育期间耕层65%～75%田间持水量,可促进棉花根系向下生长,有利于实现膜下滴灌棉花节水高产高效生产.     

Gravitropic response growth angle and vertical distribution of roots of wheat (Triticum aestivum L.)

Recent work on root distribution, growth angles and gravitropic responses in Japanese cultivars of winter wheat are reviewed. Vertical distribution of roots, which influences the environmental stress tolerance of plants, was observed in the 12 Japanese cultivars in the field. The root depth index (RDI: the depth at which 50% of the root length has been reached) differed among the cultivars at the stem elongation stage. Since the RDI was closely related to the growth angle of seminal roots obtained in a pot experiment, it was assumed that growth angle is useful for predicting vertical root distribution among wheat genotypes. Gravitropic responses of the primary seminal root of 133 Japanese wheat cultivars assessed by measuring the growth angle in agar medium, were larger in the northern Japanese cultivars and smaller in the southern ones. It was also found that the geographical variation resulted from the wheat breeding process, i.e. genotypes with limited gravitropic responses of roots had been selected in the southern part of Japan where excessive soil moisture is one of the most serious problems.     

Transfer of water from roots into dry soil and the effect on wheat water relations and growth

This investigation was performed to study the effect on plant water relations and growth when some of roots grow into dry soil. Common spring water (Triticum aestivum) plants were grown from seed in soil in 1.2 m long PVC (polyvinyl chloride) tubes. Some of the tubes had a PVC partition along their center so that plants developed a split root system (SPR). Part of the roots grew in fully irrigated soil on one side of the partition while the rest of the roots grew into a very dry (-4.1 MPa) soil on the other side of the partition. Split root plants were compared with plants grown from emergence on stored soil moisture (STOR) and with plants that were fully irrigated as needed (IRR). The experiment was duplicated over two temperature regimes (10°/20°C and 15°/25°C, night/day temperatures) in growth chambers. Data were collected on root dry matter distribution, soil moisture status, midday leaf water potential (LWP), leaf relative water content (RWC) and parameters of plant growth and yield.Some roots were found in the dry side of SPR already at 21 DAE (days after emergence) at a soil depth of 15 to 25 cm. Soil water potential around these roots was -0.7 to -1.0 MPa at midday, as compared with the initial value of -4.1 MPa. Therefore, water apparently flowed from the plant into the dry soil, probably during the night. Despite having most of their roots (around 2/3 of the total) in wet soil, SPR plants developed severe plant water stress, even in comparison with STOR plants. Already at 21 DAE, SPR plants had a LWP of -1.5 to -2.0 MPa, while IRR and STOR had a LWP of -0.5 MPa or higher. As a consequence of their greater plant water stress, SPR as compared with IRR plants were lower in tiller number, ear number, shoot dry matter, root dry matter, total biomass, plant height and grain yield and had more epicuticular wax on their leaves.It was concluded that the exposure of a relatively small part of a plant root system to a dry soil may result in a plant-to-soil water potential gradient which may cause severe plant water stress, leading to reduced plant growth and yield.     

Touch modulates gravity sensing to regulate the growth of primary roots of Arabidopsis thaliana

Plants must sense and respond to diverse stimuli to optimize the architecture of their root system for water and nutrient scavenging and anchorage. We have therefore analyzed how information from two of these stimuli, touch and gravity, are integrated to direct root growth. In Arabidopsis thaliana, touch stimulation provided by a glass barrier placed across the direction of growth caused the root to form a step-like growth habit with bends forming in the central and later the distal elongation zones. This response led to the main root axis growing parallel to, but not touching the obstacle, whilst the root cap maintained contact with the barrier. Removal of the graviperceptive columella cells of the root cap using laser ablation reduced the bending response of the distal elongation zone. Similarly, although the roots of the gravisensing impaired pgm1-1 mutant grew along the barrier at the same average angle as wild-type, this angle became more variable with time. These observations imply a constant gravitropic re-setting of the root tip response to touch stimulation from the barrier. In wild-type plants, transient touch stimulation of root cap cells, but not other regions of the root, inhibited both subsequent gravitropic growth and amyloplast sedimentation in the columella. Taken together, these results suggest that the cells of the root cap sense touch stimuli and their subsequent signaling acts on the columella cells to modulate their graviresponse. This interaction of touch and gravity signaling would then direct root growth to avoid obstacles in the soil while generally maintaining downward growth.     

Adaptive longitudinal growth of first-order lateral roots of a woody species (Spartium junceum) to slope and different soil conditions—upward growth of surface roots

First-order lateral roots originating in the upper part of the taproot of a woody species, usually termed surface roots, grow close beneath the soil surface, even on irregular or sloping ground. In slope condition, in fact, the surface roots can assume upward as well as downward growth. Existing knowledge on the controls over root direction does not fully explain these field observations.

Two different soil types and sloping conditions were selected in field condition to explore the behaviour of the surface roots in the woody species Spartium junceum L. The root system 3D architecture was measured with a 3D digitizer and the angle of growth (0° = vertically downwards) and the radial direction (0° = horizontally downslope or northwards) of all root segments measured.

Surface roots were more numerous in clay soil than in loam soil, independently from the slope inclination. They had initial angles larger than 90°, i.e. they grew upwards only in clay soil. The subsequent angles of growth maintained this value only in steep-slope condition, showing a clear soil type x slope inclination interaction. The initial angle of all first-order lateral roots decreased linearly with depth of origin on the taproot always in relation to the soil type, with this relationship being stronger in clay soil.

These findings showed that the liminal angle (the preferred angle of growth) of surface roots was mainly affected by the soil type rather than the soil surface inclination. Thus, upward growth must stand in the plasticity of the plagiotropic response of these secondary laterals rather than in a strong internal control.     

The gradient between pre-dawn rhizoplane and bulk soil matric potentials, and its relation to the pre-dawn root and leaf water potentials of four species

Uptake of soil water by plants may result in significant gradients between bulk soil and soil in the vicinity of roots. Few experimental studies of water potential gradients in close proximity to roots, and no studies on the relationship of water potential gradients to the root and leaf water potentials, have been conducted. The occurrence and importance of pre-dawn gradients in the soil and their relation to the pre-dawn root and leaf water potentials were investigated with seedlings of four species. Pre-germinated seeds were grown without watering for 7 and lid in a silt loam soil with initial soil matric potentials of -0.02, -0.1 and -0.22 MPa. Significant gradients, independent of the species, were observed only at pre-dawn soil matric potentials lower than -0.25 MPa; the initial soil matric potentials were -0.1 MPa. At an initial bulk soil matric potential of -0.22 MPa, a steep gradient between bulk and rhizoplane soil was observed after 7 d for maize (Zea mays L. cv. Issa) and sunflower (Helianthus annuus L. cv. Nanus), in contrast to barley (Hordeum vulgare L. cv. Athos) and wheat (Triticum aestivum L. cv. Kolibri). Pre-dawn root water potentials were usually about the same as the bulk soil matric potential and were higher than the rhizoplane soil matric potential. Pre-dawn root and leaf water potentials tended to be much higher than rhizoplane soil matric potentials when the latter were lower than -0.5 MPa. It is concluded that plants tend to become equilibrated overnight with the wetter bulk soil or with wetter zones in the bulk soil. Plants can thus circumvent negative effects of localized steep pre-dawn soil matric potential gradients. This may be of considerable importance for water uptake and growth in drying soil.     

The configuration of the seminal roots of Triticum aestivum L. (Poaceae)

The seminal root system of wheat (Triticum aestivum L.) is composed of the primary seminal root, the first pair of seminal roots, and the second pair of seminal roots, which are known to grow in different directions. The direction of root growth, which can be expressed by theta (the angle between the root and the plumb line) and phi (the angle between the root and a vertical plane including the primary seminal root), was studied with special attention to the latter. It was measured on seedlings grown in a small hemispherical soil-filled mesh basket. There were varietal differences in the phi of the first pair of roots (phi f) and in the phi of the second pair of roots (phi s). phi f and phi s were significantly correlated. The mean distance (MD), a measure to evaluate the efficiency of root spacing, was correlated with the difference between phi f and phi s. Neither experimentally applied low soil water potential nor the excision of the primary seminal root affected phi. When the grain was sown vertically with the tip of the embryo pointing downwards, it was found that the growth movement into a direction different from the plumb line and phi s was greatly modified. It is suggested that certain internal mechanisms, possibly involving gravitropic reactions, are operating to control the direction of root growth. The significance of root growth direction at the seedling stage is discussed.     

Effect of phosphorus availability on basal root shallowness in common bean

Root gravitropism may be an important element of plant response to phosphorus availability because it determines root foraging in fertile topsoil horizons, and thereby phosphorus acquisition. In this study we seek to test this hypothesis in both two dimensional paper growth pouch and three-dimensional solid media of sand and soil cultures. Five common bean (Phaseolus vulgaris L.) genotypes with contrasting adaptation to low phosphorus availability were evaluated in growth pouches over 6 days of growth, and in sand culture and soil culture over 4 weeks of growth. In all three media, phosphorus availability regulated the gravitropic response of basal roots in a genotype-dependent manner. In pouches, sand, and soil, the phosphorus-inefficient genotype DOR 364 had deeper roots with phosphorus stress, whereas the phosphorus-efficient genotype G19833 responded to phosphorus stress by producing shallower roots. Genotypes were most responsive to phosphorus stress in sand culture, where relative root allocation to the 0–3- and 3–6-cm horizons increased 50% with phosphorus stress, and varied 300% (3–6 cm) to 500% (0–3 cm) among genotypes. Our results indicate that (1) phosphorus availability regulates root gravitropic growth in both paper and solid media, (2) responses observed in young seedlings continue throughout vegetative growth, (3) the response of root gravitropism to phosphorus availability varies among genotypes, and (4) genotypic adaptation to low phosphorus availability is correlated with the ability to allocate roots to shallow soil horizons under phosphorus stress.     

低温潜沼性逆境下不同耐受能力水稻根系动态变化

水稻根系生长分布与地上部生长及产量形成密切相关,定量研究低温潜沼性逆境下水稻根系分布特征对改进稻田管理方式及耐潜品种的选育等具有重要意义。以4个耐潜等级的8个水稻品种为实验材料,在水稻栽培环境模拟系统中连续栽植3年(2012—2014年),研究水稻根系在低温潜沼性逆境下的动态变化规律和分布规律,明确低温潜沼性逆境下水稻根系变化与产量构成的关系。结果表明:1)低温潜沼性逆境下水稻根系长度和数量的M值(绝对变化率)均大于0,P值(相对变化率)均小于0。表明,低温潜沼性逆境下水稻根系仍有一定增长,但与非低温潜沼性逆境相比,根系长度和数量均显著降低,降低程度因品种耐潜等级的高低而不同,耐潜指数高的品种(泸优5号和C优130)降低程度显著低于耐潜指数低的品种(川谷优204和协优027)。2)低温潜沼性逆境对0—10 cm土层根系的影响在5月底6月初最大;对10—30 cm土层根系的影响在6月底最大;对30—50 cm土层根系的影响在6月30日前后最大。3)低温潜沼性逆境下,水稻孕穗-抽穗期根系长度和数量与穗粒数、实粒数、结实率、产量之间呈显著正相关;与单株有效穗数和千粒重呈显著负相关,根系长度和数量的M值与P值,尤其是白根长度和数量的M值与P值可作为一个水稻耐潜能力评价指标进行考虑。4)低温潜沼性逆境下,不同土层水稻根系与产量构成因素间的相关性不同。0—10 cm土层根系长度和数量与有效分蘖数呈显著负相关;10—20 cm土层白根长度和数量与结实率呈显著正相关;30—50 cm土层根系长度和数量与着粒数呈显著正相关。     

Influence of soil water deficits on root growth of cotton seedlings

Summary Cotton (Gossypium hirsutum L. cv. H14) seedlings were raised in soil of differing soil water content in specially designed pots in which the roots had access to freely available water and nutrients located 2.5 cm below the base of the soil core. The time for root emergence from the soil core and the rate of root growth were measured daily from sowing to harvest. The root and shoot dry weight and leaf water potential were measured at the final harvest 16 days after sowing. As soil water content decreased, the root emerged from the soil earlier and the initial rate of root elongation was faster. In spite of the availability of freely available water, the plants in the soil at low water contents had significantly lower leaf water potentials than those in soil at high water contents. The root: shoot ratio increased as the soil water content decreased. This arose from an absolute increase in root weight, with shoot weight not being significantly affected.     

Simulation of the effect of root distribution on hydraulic redistribution in a desert riparian forest

Hydraulic redistribution (HR) of roots plays an important role in the water relations of desert riparian plants. In order to estimate the effect of vertical root distribution on the HR process of Populus euphratica Oliv. during the entire growth season, we performed simulation and scenario analyses based on the observed soil water potential and root distribution data. The results showed that our simulation model achieved a good accuracy. The initial value of soil water content could significantly affect the simulated soil water content at soil depths of >90 cm, but had only limited effect on soil water content in the 0- to 90-cm soil layers. Scenario analysis revealed that with increase in root distribution depth, the HR process extended from the upper and middle soil layers downward toward the middle and deeper soil layers: the deeper the root distribution, the more likely it was to trigger the HR process in deep soil layers. However, a deeper rooting system led to a decrease in the total amount of hydraulically redistributed water over the entire soil column. Redistributed water also significantly increased the soil water depletion and the soil water storage. However, the effects of redistributed water (HR vs. without HR) on water depletion and soil water storage were reduced with the deepening of root distribution. These results indicate that HR can obviously affect the moisture of the upper soil layers, while vertical root distribution significantly changes the spatial and quantitative characteristics of HR within soil columns.     

根区交替控制灌溉条件下玉米根系吸水规律

为了阐明根区交替控制灌溉(CRDAI)条件下玉米根系吸水规律,通过田间试验,在沟灌垄植模式下采用根区交替控制灌溉研究玉米根区不同点位(沟位、坡位和垄位)的根长密度(RLD)及根系吸水动态。研究表明,根区土壤水分的干湿交替引起玉米RLD的空间动态变化,在垄位两侧不对称分布,并存在层间差异;土壤水分和RLD是根区交替控制灌溉下根系吸水速率的主要限制因素。在同一土层,根系吸水贡献率以垄位最大,沟位最低;玉米营养生长阶段,10—30 cm土层的根系吸水速率最大;玉米生殖生长阶段,20—70 cm为根系吸水速率最大的土层,根系吸水贡献率为43.21%—55.48%。研究阐明了交替控制灌溉下根系吸水与土壤水分、RLD间相互作用的动态规律,对控制灌溉下水分调控机理研究具有理论意义。     

用Minirhizotrons观测柠条根系生长动态

 Minirhizotrons是一种非破坏性、定点直接观察和研究植物根系的新方法。该文介绍了用Minirhizotrons测定植物根系的方法,并同根钻取原状土样法进行了比较;探讨了根系生长动态同土壤含水量间的关系。试验于2004年植物生长季在沙坡头沙漠试验研究站的水分平衡观测场的人工柠条(Caragana korshinskii)林进行,结果表明：Minirhizotrons 管埋入土壤后需要10个月时间允许柠条根系在其周围定居,其观测图片中的根系代表了管子周围2.6 mm土层的根系。柠条根系生长动态和土壤水分变化相关,含水量的升高导致根系的大量繁殖,而根系吸水及蒸发散又导致含水量的减少;在2004年植物生长季,土壤水分和根系的这种相互作用出现了两次,但根系生长高峰比土壤含水量高峰滞后20 d左右。     

疏花水柏枝幼苗生物量与构件对模拟土壤地下水位变化的响应

通过模拟不同地下水位的方法,对疏花水柏枝（Myricaria laxiflora（Franch.）P.Y.Zhang et Y.J.Zhang）一年生幼苗在不同条件下地上与地下部生物量及构件的变化进行测定,分析幼苗生长对地下水位变化的响应。结果显示：随着地下水位的降低,疏花水柏枝幼苗的生长特征指标均呈先增加后减少的趋势,其中地上、地下部分生物量的最高值分别为0.0438、0.0100 g,最低值分别为0.0177、0.0026 g。幼苗地上部生物量在-10 cm处理水平最高;地下部生物量在-15 cm处理水平最高。幼苗直径、根表面积、株高、主根长度、根体积、一级枝数、二级枝数等指标也分别在-10 cm或-15 cm处理水平达到最高值。疏花水柏枝幼苗主要构件的生长与地下水位的变化存在显著相关性。主成分分析结果表明,幼苗的地下部分更容易受到土壤地下水位变化的影响,幼苗性状症候群随地下水位的变化而发生移动,说明该物种幼苗在不同地下水位时的生长投资策略具有较大差异。     

轮耕对关中一年两熟区土壤物理性状和冬小麦根系生长的影响

针对关中地区土壤连续单一耕作存在的主要问题,进行了土壤轮耕效应研究。2009年至2012年在关中一年两熟区采用连续4a旋耕(RT)、翻耕-免耕-翻耕-免耕(PNT)和深松-免耕-深松-免耕(SNT)3种耕作处理,对土壤容重、紧实度及小麦根系生长进行了研究。结果表明,与试验前相比,夏玉米收获后(2013年10月)两种轮耕处理显著(P0.05)降低了0—10、10—20 cm土壤容重,旋耕处理在0—10 cm处差异不显著,而10—20 cm土壤容重显著增大;与旋耕处理相比,两种轮耕处理0—10、10—20 cm土壤容重在第4季冬小麦整个生育期内变异系数较小,土壤紧实度较低,且改善效果在冬小麦生育中后期10—20 cm土层体现更为显著;旋耕处理0—10、10—20 cm土壤紧实度与含水量均呈显著负相关,相关系数分别为-0.89、-0.85,两种轮耕处理相关性不显著;0—40 cm土层根重密度和根系活力表现为:两种轮耕处理连年旋耕。可见,长期旋耕后进行轮耕(免耕与翻耕、深松)有利于改善土壤物理状况,促进作物根系生长。